Abstract
The aim of this paper is to describe a new species of the genus Loxosceles Heineken & Lowe, 1832 from Morocco, Loxosceles imazighen sp. n., and to describe for the first time a female of Loxoxceles mrazig Ribera & Planas, 2009 from Tunisia. Both species live in xeric and desert environments and are located in southern Atlas Range. Molecular phylogenetic analyses, using mitochondrial (cox1, 16S) and nuclear (H3, 28S) markers, revel that these species are closely related and that they constitute a separate evolutionary lineage of L. rufescens (Dufour, 1820) and of the set of endemic species of the Canary Islands. L. imazighen sp. n. differs from L. mrazig, the closest species morphologically and geographically, in the shapes and proportions of the male palpal tibia and the shapes and dispositions of the female seminal receptacles. In addition, L. mrazig females show morphological variability in their genitalia, mainly in the inner and outer lobes. Although that variability cannot be associated with different populations, since it also appears within individual populations, and is not related to genetic or geographic distances.
References
Arnedo, M.A., Oromí, P. & Ribera, C. (2001) Radiation of the spider genus Dysdera (Araneae, Dysderidae) in the Canary Islands: Cladistic assessment based on multiple data sets. Cladistics, 17, 313–353. https://doi.org/10.1111/j.1096-0031.2001.tb00129.x
Bond, J.E. & Hedin, M.C. (2006) A total evidence assessment of the phylogeny of the diverse North American trapdoor spider subfamily Euctenizinae (Araneae, Mygalomorphae, Cyrtaucheniidae). Molecular Phylogenetics and Evolution, 4, 70–85. https://doi.org/10.1016/j.ympev.2006.04.026
Brignoli, P.M. (1976) Beiträge zur Kenntnis der Scytodidae (Araneae). Revue Suisse de Zoologie, 83 (1), 125–191.
Colgan, D.J., McLauchlan, A., Wilson, G.D.F., Livingston, S.P., Edgecombe, G.D., Macaranas, J., Cassis, G. & Gray, M.R. (1998) Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology, 46, 419–437. https://doi.org/10.1071/ZO98048
Duncan, R. P., Rynerson, M.R., Ribera, C. & Binford, G.J. (2010) Diversity of Loxosceles spiders in Northwestern Africa and molecular support for cryptic species in the Loxosceles rufescens lineage. Molecular Phylogenetics and Evolution, 55 (1), 234–248. https://doi.org/10.1016/j.ympev.2009.11.026
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular marine biology and biotechnology, 3 (5), 294–299. [PMID: 7881515]
Gertsch, W.J. (1967) The spider genus Loxosceles in South America (Araneae, Scytodidae). Bulletin of the American Museum of Natural History, 136, 117–174.
Habel, J.C., Husemann, M., Schmitt, T., Zachos, F.E., Honnen, A-C., Petersen, B., Parmakelis, A. & Stathi, L. (2012) Microallopatry caused strong diversification in Buthus scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa). PLoS ONE, 7 (2), e29403. https://doi.org/10.1371/journal.pone.0029403
Hedin, M.C. & Maddison, W.P. (2001) A combined molecular approach to phylogeny of the jumping spider subfamily Dendryphantinae (Araneae: Salticidae). Molecular Phylogenetics and Evolution, 18, 386–403. https://doi.org/10.1006/mpev.2000.0883
Huber, B.A., Pérez González, A., Astrin, J.J., Blume, C., Baptista, R. (2013) Litoporus iguassuensis Mello-Leitão, 1918 (Araneae, Pholcidae): camouflaged retreat, sexual dimorphism, female color polymorphism, intraspecific genital variation, and description of the male. Zoologischer Anzeiger, 252, 511–521. https://doi.org/10.1016/j.jcz.2012.12.001
Katoh, K.& Toh, H. (2008) Improved accuracy of multiple ncRNA alignment by incorporating structural information into a MAFFT-based framework. BMC Bioinformatics, 9, 212. https://doi.org/10.1186/1471-2105-9-212
Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., †Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28 (12), 1647–1649. https://doi.org/10.1093/bioinformatics/bts199
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874. https://doi.org/10.1093/molbev/msw054
Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29, 1695–1701. https://doi.org/10.1093/molbev/mss020
Lotz, L.N. (2017) An update on the spider genus Loxosceles (Araneae: Sicariidae) in the Afrotropical region, with description of seven new species. Zootaxa, 4341 (4) 475–494. https://doi.org/10.11646/zootaxa.4341.4.2
Mallatt, J. & Sullivan, J. (1998) 28S and 18S rDNA sequences support the monophyly of lampreys and hagfishes. Molecular Biology and Evolution, 15, 1706–1718. https://doi.org/10.1093/oxfordjournals.molbev.a025897
Massa, M., Planas, E. & Ribera, C. (2018) The Mediterranean as a melting pot: Phylogeography of Loxosceles rufescens (Sicariidae) in the Mediterranean Basin. PLoS ONE, 13 (12), 1–10. https://doi.org/10.1371/journal.pone.0210093
Millot, J. (1941) Les araignées de l’Afrique Occidentale Française - sicariides et pholcides. Mémoires de l’Académie des Sciences de l’Institut de France, 64, 1–53.
Nentwig, W., Pantini, P. & Vetter, R.S. (2017) Distribution and medical aspects of Loxosceles rufescens, one of the most invasive spiders of the world (Araneae: Sicariidae), Toxicon, 32, 19–28. https://doi.org/10.1016/j.toxicon.2017.04.007
Planas, E., Fernandez-Montraveta, C. & Ribera, C. (2013) Molecular systematics of the wolf spider genus Lycosa (Araneae: Lycosidae) in the Western Mediterranean Basin. Molecular Phylogenetics and Evolution, 67, 414–428. https://doi.org/10.1016/j.ympev.2013.02.006
Planas, E. & Ribera, C. (2014) Uncovering overlooked island diversity: colonization and diversification of the medically important spider genus Loxosceles (Arachnida: Sicariidae) on the Canary Islands. Journal of Biogeography, 41, 1255–1266. https://doi.org/10.1111/jbi.12321
Planas, E. & Ribera, C, (2015) Description of six new species of Loxosceles (Araneae: Sicariidae) endemic to the Canary Islands and the utility of DNA barcoding for their fast and accurate identification. Zoological Journal of the Linnean Society, 174 (1), 47–73. https://doi.org/ 10.1111/zoj.12226
Planas, E., Saupe, E.E., Lima-Ribeiro, M.S., Townsend, P.A. & Ribera, C. (2014) Ecological niche and phylogeography elucidate complex biogeographic patterns in Loxosceles rufescens (Araneae, Sicariidae) in the Mediterranean Basin. BMC Evolutionary Biology, 14, 195. https://doi.org/10.1186/s12862-014-0195-y
Ribera, C. & Planas, E. (2009) A new species of Loxosceles (Araneae, Sicariidae) from Tunisia. ZooKeys. 16, 217–225. https://doi.org/10.3897/zookeys.16.232
Ribera, C., Elverici, M., Kunt, K.B. & Özkütük, R.S. (2014) Typhlonesticus gocmeni sp. n., a new cave-dwelling blind spider species from the Aegean region of Turkey (Araneae, Nesticidae). ZooKeys, 419, 87–102. https://doi.org/10.3897/zookeys.419.5739
Satler, J.D., Carstens, B.C. & Hedin, M. (2013). Multilocus species delimitation in a complex of morphologically conserved trapdoor Spiders (Mygalomorphae, Antrodiaetidae, Aliatypus). Systematic Biology, 62 (6), 805–823. https://doi.org/doi:10.1093/sysbio/syt041
Silvestro, D. & Michalak, I. (2012) RaxmlGUI: A graphical front-end for RAxML. Organisms Diversity and Evolution, 12, 335–337. https://doi.org/doi.org/10.1007/s13127-011-0056-0
Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene-sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–701. https://doi.org/10.1093/aesa/87.6.651
Stamatakis, A. (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixedmodels. Bioinformatics, 22, 2688–2690. https://doi.org/10.1093/bioinformatics/btl446
Tahami, M.S., Zamani, A., Sadeghi, S. & Ribera, C. (2017) A new species of Loxosceles Heineken & Lowe, 1832 (Araneae: Sicariidae) from Iranian caves. Zootaxa, 4318 (2), 377–387. https//doi.org/10.11646/zootaxa.4318.2.10
Vetter, R.S. (2008) Spiders of the genus Loxosceles (Araneae, Sicariidae): a review of biological, medical and psychological aspects regarding envenomation. Journal of Arachnology, 36, 150–163. https//doi.org/10.1636/RSt08-06.1
WSC [World Spider Catalog] (2021) World Spider Catalog. Version 22.5. Natural History Museum Bern, Bern. Available from: http://wsc.nmbe.ch (accessed 15 September 2021)
Zamani, A., Mirshamsi, O. & Marusik, Y.M. (2021) ‘Burning violin’: the medically important spider genus Loxosceles (Araneae: Sicariidae) in Iran, Turkmenistan, and Afghanistan, with two new species. Journal of Medical Entomology, 58 (2), 666–675. https//doi.org/10.1093/jme/tjaa257